In this paper we describe a fossil fungus-Paleoblastocladia milleri ge
n. et sp. nov.-from the 400 million-year-old Early Devonian Rhynie che
rt that shares numerous features with modern zoosporic fungi placed in
the order Blastocladiales. The fungus occurs in tufts that arise from
stomata or between the cuticle and epidermis of Aglaophyton major axe
s. Thallus development begins from an irregular bipolar basal cell tha
t produces a system of intramatrical rhizoids and clavate-shaped extra
matrical, nonseptate hyphae. These hyphae develop into two types of ma
ture thalli. Sporothalli are characterized by several orders of dichot
omous branching and the production of terminal, globose zoosporangia,
as well as thick-walled, pitted resting sporangia. On separate dichoto
mously branched thalli (gametothalli) are terminal chains of two or th
ree gametangia, in which the terminal one is slightly larger. Despite
the fact that all of the reproductive organs contain either zoospores
or gametes, none show evidence of discharge papillae. The fossil fungu
s is compared with extant members of the Blastocladiales, and the pres
ence of sexual reproduction is discussed.